disentangling palaeodiversity signals from a biased...

Disentangling palaeodiversity signals from a biased sedimentary

record: an example from the Early to Middle Miocene of Central

Paratethys Sea

MARTIN ZUSCHIN1*, MATHIAS HARZHAUSER2 & OLEG MANDIC2

1University of Vienna, Department of Palaeontology, Althanstrasse 14, A-1090 Vienna, Austria2Natural History Museum Vienna, Burgring 7, A-1010 Vienna, Austria

*Corresponding author (e-mail: [email protected])

Abstract: Changes in molluscan diversity across the 3rd order sequence boundary from the Lowerto the Middle Miocene of the Paratethys were evaluated in the context of environmental bias.Taken at face value, quantitative data from nearshore and sublittoral shell beds suggest a transitionfrom low-diversity Karpatian (Upper Burdigalian) to highly diverse Badenian (Langhian andLower Serravallian) assemblages, but environmental affiliation of samples reveals a strongfacies shift across the sequence boundary. Ordination methods show that benthic assemblages ofthe two stages, including 4 biozones and four 3rd order depositional sequences over less thanfour million years, are developed along the same depth-related environmental gradient. Almostall samples are from highstand systems tracts, but Karpatian faunas are mostly from nearshore set-tings, and Badenian faunas are strongly dominated by sublittoral assemblages. This study empha-sizes the importance of highly resolved stratigraphic and palaeoenvironmental frameworks fordeciphering palaeodiversity patterns at regional scales and highlights the effort required to reachthe asymptote of the collector’s curve. Abundance data facilitate the recognition of ecologicalchanges in regional biota and it is suggested that in second and higher order sequences thefacies covered within systems tracts will drive observed diversity patterns.

The quality of the fossil record of biodiversity isstrongly influenced by the rock record (Holland2000; Smith 2007). The amount of sedimentaryrock preserved has strongly fluctuated over timeand is very similar to corresponding diversity pat-terns, suggesting that a major bias exists (Raup1976; Miller & Foote 1996; Smith 2001; Peters &Foote 2001; Smith & McGowan 2007; Barrettet al. 2009). Alternatively, it suggests that both therock record and diversity are driven by a commonunderlying factor, such as sea-level change (Peters2005, 2006), a signal that can be regionally obscuredat tectonically active margins (Crampton et al.2003). The change in the proportion of onshore tooffshore sediments preserved in the record, how-ever, is probably as important as changes in thevolume of rock preserved (Smith et al. 2001;Crampton et al. 2003, 2006). Although global data-sets are comparatively robust to such biases (e.g.Marx & Uhen 2010), sequence stratigraphical archi-tecture undoubtedly controls patterns of faunalchange on a local and regional scale (Bulot 1993;Brett 1995, 1998; Patzkowsky & Holland 1999;Smith et al. 2001; Smith 2001). Specifically, mostchanges in first and last occurrences of species,and widespread changes in species abundance andbiofacies, occur at sequence boundaries and at major

transgressive surfaces (Holland 1995, 1999, 2000).It is therefore important to evaluate stage-levelchanges in taxonomic diversity, at the temporalscale of millions of years, in the context of rockvolume- and environmental bias to ensure thatthese changes are not simply driven by sequencearchitecture (Smith 2001).

The present study focuses on diversities of tworegional Miocene stages of the Paratethys, an epi-continental sea whose history is closely linked tothe Alpine orogeny and that covered vast parts ofCentral and Eastern Europe (Rogl 1998, 1999)(Fig. 1). Standing diversity of the Central Paratethysindicates a strong increase in species richness atthe boundary from the Karpatian (Upper Burdiga-lian) to the Badenian (Langhian and Lower Serra-vallian), which is interpreted as a major faunalturnover associated with the Langhian transgression(Harzhauser et al. 2003; Harzhauser & Piller 2007).Based on a comprehensive echinoderm dataset,however, it has been suggested that the low diver-sity of the Karpatian was rather caused by non-preservation of suitable habitats (Kroh 2007). Inthis study we use a species abundance dataset ofbenthic molluscs to evaluate the influence of envi-ronmental bias on the faunal change. Previous mol-luscan species lists from the area are not useful for

From: McGowan, A. J. & Smith, A. B. (eds) Comparing the Geological and Fossil Records: Implicationsfor Biodiversity Studies. Geological Society, London, Special Publications, 358, 123–139.DOI: 10.1144/SP358.9 0305-8719/11/$15.00 # The Geological Society of London 2011.

this purpose because they are biased in favour oflarger shells and biostratigraphically and palaeogeo-graphically useful species, but stable temporal andspatial patterns of diversity can only be decipheredusing large bulk samples from extensive fieldwork (Koch 1978; Kosnik 2005).

The Vienna Basin and adjacent basins have nowbeen systematically studied for almost two centuriesfor molluscs and other invertebrates. Based on thepublication of a visiting French geologist (Prevost1820), these basins were key areas for the foun-dation of the concept of the Tertiary in the early19th century (Rudwick 2005, pp. 546–549; Vavra2010). The stratigraphy of the Central Paratethysis comparatively well studied (for review seePiller et al. 2007) and the taxonomic compositionof the Central Paratethys molluscan fauna verywell known (e.g. Schultz 2001, 2003, 2005). Compi-lations on the standing diversity of Neogene stageswere published recently (Harzhauser et al. 2003,Harzhauser & Piller 2007). With respect to species-abundance patterns, it has been shown that at thescale of outcrops, shell beds, and samples mostspecies are rare and diversity is patchy (Zuschinet al. 2004a, 2006), a pattern that is also evidentfor the total assemblage studied here. Diversity isinfluenced by taphonomic processes, for example

by size sorting during tempestitic transport(Zuschin et al. 2005). Finally, it has been suggestedthat diversities of the marine Paratethys are lowerthan those of contemporary adjacent basins becausediversity curves have rather gentle slopes whencompared with such curves from the MioceneBoreal bioprovince (Kowalewski et al. 2002).

So far, however, studies dealing with potentialbiases of the raw diversities, including samplingefficiency, stage duration, fossil preservation orrock record bias, are scarce for the Paratethys(Kroh 2007). Studies on the quantitative compo-sition of fossil molluscan lagerstatten have onlybeen performed during the last few years (see refer-ences in Table 1). The present contribution is thefirst attempt to link this information to better under-stand one of the strongest diversity turnovers in theCentral Paratethys, the transition from the Karpatianto the Badenian (Harzhauser et al. 2003; Harzhauser& Piller 2007).

Geological setting

The Paratethys was an epicontinental sea rangingfrom the French/Swiss border region in the westto the Transcaspian area (east of Lake Aral in

Fig. 1. Karpatian and Badenian palaeogeography of the Central Paratethys (modified after Rogl (1998) and Kovacet al. (2004a, 2007)) with approximate positions of studied localities. Karpatian (i.e. Upper Burdigalian) localitiesinclude: 1 ¼ Laa a.d. Thaya; 2 ¼ Kleinebersdorf; 3 ¼ Neudorf bei Staatz; 4 ¼ Korneuburg SPK. Badenian (i.e.Langhian and Lower Serravalian) localities include: 5 ¼ Grund; 6 ¼ Immendorf; 7 ¼ Gainfarn; 8 ¼ St. Veit;9 ¼ Borsky Mikulas, 10 ¼ Niederleis.

M. ZUSCHIN ET AL.124

Table 1. Basic data of the studied assemblages

Locality Section Stageinternational

Stageregional

Biozone benthicforaminifers

Formation Sequencestratigraphy(3rd order)

Systemstract

Age Geographical position No. ofshellbeds

No. ofsamples

References

Latitude Longitude

Laa a.d. Thaya WienerbergerAG

Burdigalian Karpatian Uvigerinagraciliformis

NovyPrerovFm

Tb.2.2 Late HST 16.5 48843′07′′ 16824′57′′ 1 4 Unpublisheddata

Kleinebersdorf KleinebersdorfSandpitLehner


KorneuburgFm

Tb.2.2 Late HST 16.5 48829′37′′ 16823′44′′ 1 3 Zuschin et al.2004a

KleinebersdorfSandpitWohlmuth


KorneuburgFm

Tb.2.2 Late HST 16.5 48829′42′′ 16823′48′′ 1 3 Zuschin et al.2004a

Korneuburg KorneuburgSPK


KorneuburgFm


Neudorf beiStaatz


NovyPrerovFm


Grund Langhian Badenian Lower LagenidaeZone

Grund Fm Tb.2.3 Early HST 15 48838′18′′ 16803′48′′ 5 5 Zuschin et al.2004b, 2005

Immendorf Langhian Badenian Lower LagenidaeZone

Grund Fm Tb.2.3 Early HST 15 48839′00′′ 16807′49′′ 5 25 Zuschin et al.2006

Niederleis NiederleisBuschberg

Langhian Badenian Lower LagenidaeZone

Lanzhot Fm Tb.2.3 Early HST 15 48833′48′′ 16824′17′′ 4 4 Mandic et al.2002

NiederleisBahnhof

Langhian Badenian Lower LagenidaeZone

Lanzhot Fm Tb.2.3 Early HST 15 48832′25′′ 16824′39′′ 5 5 Mandic et al.2002

St. Veit a.dTriesting

Langhian Badenian Upper LagenidaeZone

Lanzhot Fm Tb.2.3 Late HST 14.5 47855′55′′ 16808′53′′ 9 9 Unpublisheddata

Gainfarn Gainfarn 1 Langhian Badenian Upper LagenidaeZone

Lanzhot Fm Tb.2.3 Late HST 14.5 47856′45′′ 16810′59′′ 7 8 Zuschin et al.2007

Gainfarn 2 Langhian Badenian Upper LagenidaeZone

Jakubov Fm Tb.2.4 TST 14 47856′40′′ 16810′57′′ 14 14 Zuschin et al.2007

Borsky Mikulas Serravallian Badenian Bolivina/BuliminaZone

StudienkaFm

Tb.2.5 Early HST 13 48836′20′′ 17811′57′′ 3 17 Svagrovsky1981;Unpublisheddata

PA

LA

EO

DIV

ER

SIT

YF

RO

MA

BIA

SE

DS

ED

IME

NT

AR

YR

EC

OR

D125

Kazakhstan) in the east. Its development startedduring the Late Eocene to Oligocene and wasstrongly linked to the alpine orogeny. It was separ-ated from the Mediterranean by the newly formedland masses of the Alps, Dinarides, Hellenides,and the Anatolian Massif. Afterwards, it experi-enced a complex history of connection and discon-nection with the Mediterranean Sea (Rogl 1998,1999; Popov et al. 2004). The present study focuseson shell beds of the Vienna Basin and the NorthAlpine Foreland Basin; in terms of palaeogeogra-phy, they were part of the Central Paratethys,which ranged from southern Germany in the westto the Carpathian Foredeep, Ukraine in the east,and from Bulgaria in the south to Poland in thenorth (Fig. 1). Due to the complex geodynamichistory, a regional chronostratigraphic stage system(Fig. 2) is used in the Central Paratethys. The twostages of interest here are the Karpatian and theBadenian. The Karpatian stage is characterized bya strong tectonic reorganization in the CentralParatethys area, leading to a change from west–east trending basins towards rift and intra-mountainbasins (Rogl & Steininger 1983; Rogl 1998; Kovacet al. 2004b). Associated with this geodynamicimpact is the abrupt, discordant progradation ofupper Karpatian fossiliferous estuarine to shallow

marine deposits over macrofossil-poor lower Karpa-tian offshore clays in the North Alpine ForelandBasin and in the Carpathian Foredeep (Adameket al. 2003). The climate was subtropical withwarm and wet summers and rather dry winters(Harzhauser et al. 2002; Kern et al. 2010). Theearly Middle Miocene is marked by a widespreadmarine transgression following a major drop in sea-level at the Burdigalian/Langhian transition (Haqet al. 1988; Hardenbol et al. 1998). The regressionwas intensified by regional tectonic movements,the so-called Styrian phase (Stille 1924; Roglet al. 2006). Sediments of the Langhian transgres-sion are commonly eroded or reduced in thicknessat the basin borders, with continuous sedimentationoccurring only in bathyal settings of the basincentres (Hohenegger et al. 2009). In shallow-marineenvironments of the Vienna Basin, erosion of upto 400 m took place between the youngest pre-served Karpatian and the oldest preserved Badeniansediments (Strauss et al. 2006). Due to the tectonicreorganization, however, a broad connectionopened between the Mediterranean Sea and theParatethys during the Langhian transgression,through which free faunal exchange occurred(Rogl 1998; Studencka et al. 1998; Harzhauseret al. 2002; Harzhauser & Piller 2007). The rising

Fig. 2. Stratigraphic details for the studied sections and standing diversity of Karpatian and Badenian gastropodscompiled from regional species lists and monographs (after Harzhauser & Piller 2007). The sections belong tosix formations and four 3rd order sequence stratigraphic cycles and are all, except Gainfarn 2, from early or late HSTs(cf. Table 1). EBBE ¼ Early Badenian Build-up Event.


sea-level and the Middle Miocene climatic optimumpotentially strongly influenced marine life in theCentral Paratethys (Harzhauser et al. 2003). In con-trast to the Karpatian, the Badenian stage is charac-terized by highly fossiliferous offshore sands andpelites, and by carbonate platforms (corallinaceanlimestones and coral carpets). Several fossilgroups increase strongly in diversity at the onsetof the Badenian (Fig. 2). This event has been expli-citly worked out for gastropods, with 505 taxahaving their first occurrences (FOs), and for forami-nifers, with FOs of 82 taxa (Harzhauser & Piller2007). These authors dubbed this event as ‘EarlyBadenian Build-up Event’ (EBBE).

Material and methods

We studied benthic molluscs from 10 localities fromthe Karpatian (Upper Burdigalian) to the Badenian(Langhian and Lower Serravallian), covering allavailable fossil lagerstatten in the Vienna Basinand the North Alpine Foreland Basin that wereamenable to bulk sampling (Fig. 3, Table 1). Allsamples are from siliciclastic pelitic, sandy andgravelly sediments, are characterized by aragoniteand calcite preservation and were sieved through a1 mm mesh. Detailed palaeoecological and tapho-nomical studies have been published on some ofthe sections (see references in Table 1). The shellbeds of the respective localities were depositedbetween 16.5 and 12.7 Ma and belong to six for-mations, four 3rd order sequence stratigraphiccycles (Tb.2.2 to Tb.2.5 of Hardenbol et al. 1998),

and are mostly part of highstand systems tracts(HST); only one section belongs to a transgressivesystems tract (TST) (Strauss et al. 2006). All fossi-liferous Karpatian assemblages belong to a singleregional benthic foraminifera biozone, and thestudied Badenian assemblages to three such bio-zones (Table 1) (Uvigerina graciliformis zone,Lower and Upper Lagenidae zones and Bolivina/Bulimina zone; Grill 1943; Steininger et al. 1978).The faunal transition from the Karpatian to theBadenian is studied at the level of stages and bio-zones. For the purpose of this study, samples areenvironmentally assigned to the intertidal to veryshallow sublittoral (,1 m water depth), termed asnearshore for the rest of the paper, and to thedeeper sublittoral (few metres to several tens ofmetres of water depth). Palaeoenvironmental des-ignations of samples were based on palaeogeo-graphical positions of localities and actualisticenvironmental requirements of dominant molluscantaxa. Independent data from foraminifera confirmour assignments and suggest a total range of deposi-tional water depths from intertidal to several tensof metres (pers. comm. Holger Gebhart, PatrickGrunert, Johann Hohenegger & Fred Rogl, 2009).Logarithmic scale rank abundance plots of familylevel data were used to compare community organ-ization between stages and the data were fit to geo-metric series, log-series, broken stick andlog-normal abundance models using the programPAST (Hammer et al. 2001). Species accumulationcurves were computed to compare species richnessbetween stages, biozones and environments usingthe program Estimates with 50 sample order

Fig. 3. Map of sample localities in Austria and Slovakia. Karpatian (i.e. Upper Burdigalian) localities include: 1 ¼ Laaa.d. Thaya; 2 ¼ Kleinebersdorf; 3 ¼ Neudorf bei Staatz; 4 ¼ Korneuburg SPK. Badenian (i.e. Langhian and LowerSerravalian) localities include: 5 ¼ Grund; 6 ¼ Immendorf; 7 ¼ Gainfarn; 8 ¼ St. Veit; 9 ¼ Borsky Mikulas,10 ¼ Niederleis (modified after Sawyer & Zuschin 2011).

PALAEODIVERSITY FROM A BIASED SEDIMENTARY RECORD 127

randomizations without replacement. (Colwell2009). Diversity was measured as species richnessand as evenness, which is based on the proportionalabundance of species (for a review see Magurran2004). To compensate for sampling effects inspecies richness we used Margalef’s diversityindex. The Simpson index, which is affected bythe 2–3 most abundant species, and the Shannonindex, which is more strongly affected by speciesin the middle of the rank sequence of species,were used as measures of evenness (see Gray 2000for discussion). All indices were calculated usingthe program PAST (Hammer et al. 2001). The Mar-galef index was calculated with the equation

DMg = (S − 1)/ ln N

where S ¼ the total number of species and N ¼ thetotal number of individuals. The Simpson index isexpressed as 1 2 D and was calculated with theequation

D =∑S

i=1

ni(ni − 1)

N(N − 1)

where S ¼ the total number of species, ni ¼ thenumber of individuals in the ith species andN ¼ the total number of individuals. The Shannonindex was calculated with the equation

H = −∑S

i=1

pi ln pi

where S ¼ the total number of species, and pi ¼ theproportion of individuals found in the ith species.Species richness, the Simpson index and the

Shannon index were chosen because they are themost commonly employed measures of diversity(Lande 1996). It should be mentioned, however,that the underlying statistical distribution of asample will generally influence the constancy ofevenness measures and that the Shannon index isparticularly sensitive to sample size (Lande 1996;Magurran 2004; Buzas & Hayek 2005). Non-metricmultidimensional scaling (NMDS, Kruskal 1964)was used as an ordination method to evaluate thepresence of environmental gradients in the datasetand was performed using the software packagePRIMER (Clarke & Warwick 1994). Surfaceoutcrop areas and their environmental affiliation ofthe Karpatian and Badenian in Austria are adaptedfrom Kroh (2007) and were calculated from digital1:200 000 scale map series of the GeologicalSurvey of Austria for the Burgenland (Pascheret al. 2000) and Lower Austria and Vienna (Schna-bel 2002).

Results

Sampling intensity was very high (213 samples,yielding 494 species from .49 000 shells), but thespecies accumulation curve for the total assemblagedoes not level off (Fig. 4). The number of families,genera and species, however, is significantly higherfor Badenian than for the Karpatian assemblages(Fig. 5a). While in the Karpatian sampling intensitywas sufficient to cover diversity at all hierarchicallevels, for the Badenian the diversity of speciesand genera do not show a tendency to level off(Fig. 5b).

Strong differences in the abundances of domi-nant families and in the shape of the rank abundanceplot of family level data indicate environmental

Fig. 4. Species accumulation curve of the total assemblage with 95% confidence intervals. Inset: number of samples perenvironment and stage. Sampling intensity was very high but the species accumulation curve does not level off.


differences between shelly assemblages of the twostages (Fig. 6). Karpatian molluscan assemblagesare dominated by neritid and potamidid–batillariidgastropods, which indicate the prevalence of tidalflat deposits, whereas the Badenian molluscanassemblages are dominated by corbulid and ven-erid bivalves and rissoid gastropods, which all indi-cate the preponderance of sublittoral conditions(Fig. 6a). In accordance rank abundance plotssuggest higher evenness for the total Badenianassemblage (Fig. 6b) and diversity indices are sig-nificantly higher for sublittoral than for nearshoresamples in our dataset (Fig. 7). An environmentalbias may therefore explain the apparent faunal turn-over. In fact, in the Karpatian more samples derivefrom nearshore environments, whereas the Bade-nian is strongly dominated by sublittoral samples.This difference is even more pronounced when

considering biozones. In the Lower Lagenidaezone, samples are exclusively from the sublittoral;nearshore samples of the Badenian only occur inthe Upper Lagenidae zone and in the Bolivina/Buli-mina zone (Fig. 8).

At the level of stages and biozones the environ-mental affiliations of samples correlate with diver-sities, which are high wherever assemblages aredominated by samples from the sublittoral (Fig. 9).An exception is the Bulimina/Bolivina zone, butthere the sampling intensity was by far the lowest(Table 1). Species accumulation curves of environ-ments within stages and biozones are always steeperfor sublittoral than for nearshore assemblages.Differences between environments within timeslices are significant except for the Karpatian (i.e.the Uvigerina gracliformis zone) as indicated byoverlapping confidence intervals (Fig. 10). Strong

Fig. 5. Diversities in stages. (a) Average number of families, genera and species in samples with 95% confidenceintervals. (b) Taxa accumulation curves for families, genera and species of the total assemblages with 95% confidenceintervals. At all three taxonomic levels diversities differ significantly between the Karpatian and Badenian. For theBadenian the species- and genus accumulation curves do not show a tendency to level off, although sampling intensitywas very high. F ¼ families; G ¼ genera; S ¼ species.


diversity differences between sublittoral assem-blages at the level of stages and biozones indicatehabitat differences, most notably between the well-sampled Uvigerina graciliformis zone of the

Karpatian and the Lower and Upper Lagenidaezones of the Badenian (Fig. 10b). In fact, an ordina-tion of family-level data suggests the presence of adistinct water depth gradient (Fig. 11). Sublittoral

Fig. 6. Abundances of family level data for stages. (a) Percentage abundance of most important families. Karpatianassemblages are dominated by families indicative of tidal flat deposits and Badenian assemblage are dominated byfamilies indicative of sublittoral conditions. (b) Rank abundances of families suggest higher evenness in Badenianassemblages. The log normal abundance model best describes the data in both stages (Karpatian: Chi-square ¼ 5.265,p ¼ 0.7289, Badenian: Chi-square ¼ 11.73, p ¼ 0.2287); the other three tested models have very low p-values, whichimplies bad fits.


samples from the Uvigerina graciliformis zone ofthe Karpatian represent shallower environmentsthan those from the Lower and Upper Lagenidaezones. Differences between the latter can beexplained by substrate differences. Assemblagesfrom the Lower Lagenidae zone tend to be from

sandy environments and are therefore morediverse than those from the Upper Lagenidaezone, which are rather from pelitic environments.Environmental affiliation of Karpatian and Bade-nian outcrops in eastern Austria support thisfinding. In the Karpatian the importance of terres-trial, fluvial, fluvio-marine and limnic environ-ments suggests that most fossiliferous marineoutcrops are from nearshore environments. In theBadenian, in contrast, most outcrops preserve fullymarine environments (Fig. 12) (compare also Kroh2007).

Discussion

The importance of local and regional studies

The present study demonstrates that the quantitativeevaluation of bulk samples significantly improvesthe understanding of regional diversity changesat temporal scales ranging from tens of thousandsto a few million years and thereby confirms

Fig. 7. Diversity indices of samples tallied based onenvironments. Diversity indices are significantly higherfor sublittoral than for nearshore samples in our datasetand an environmental bias may therefore explain theapparent faunal turnover from the Karpatian to theBadenian.

Fig. 8. Number of nearshore and sublittoral samples instages and biozones. More Karpatian samples derivefrom nearshore environments, but the Badenian isstrongly dominated by sublittoral samples. Thisenvironmental shift is especially pronounced at the 3rdorder sequence boundary between the KarpatianUvigerina graciliformis and the Badenian LowerLagenidae zones and amplifies the impression of adiversity increase due to the Langhian transgression froma literal reading of the fossil record.

Fig. 9. Species accumulation curves with 95%confidence intervals in relation to sampled environmentsfor stages (a) and biozones (b). Diversities are highwherever assemblages are dominated by samples fromthe sublittoral. An exception is BBZ, where samplingintensity was by far the lowest. LLZ ¼ Lower Lagenidaezone; ULZ ¼ Upper Lagenidae zone; BBZ ¼ Bolivina/Bulimina zone; UgZ ¼ Uvigerina graciliformis zone.


previous authors who emphasized the importanceof rigorous, extensive sampling combined within ahighly resolved stratigraphic and palaeoenviron-mental framework for deciphering palaeodiversitypatterns (e.g. Koch 1978; Jackson et al. 1999;Kosnik 2005). Several lines of evidence suggestgreat importance of regional and local studies forthe understanding of global diversity patterns. Bio-diversity can be studied at a series of hierarchicalscales which all contribute to an understandingof its distribution in time and space (Willis &Whittaker 2002). Diversity is, however, biologically

meaningful at local scales, where ecologicalprocesses operate and at regional scales becauselocal communities receive species from a biogeo-graphically delimited metacommunity (Hubbell2001). Long-term diversity trends actually differsignificantly among major regions of the world(e.g. Miller 1997; Jablonski 1998). With respect tothe rock record there is a global diversity signaturethat relates to supercontinent cycles, but on shortertime-scales regional processes are more importantand, due to heavy sampling bias, the European andNorth American data sets drive these patterns(McGowan & Smith 2008). Correspondingly, fossilfirst and last occurrences are dominated by recordsfrom these two continents (Kidwell & Holland2002) and the Cenozoic tropics are undersampledbecause Europe and North America had largelymoved out of the tropics by Cenozoic time (Jackson& Johnson 2001). McGowan & Smith (2008) there-fore suggest focusing on the construction of regionaldata sets within tectonically and sedimentologicallymeaningful frameworks. Such regional diversitystudies can typically be performed at low taxonomic

Fig. 10. Species accumulation curves with 95%confidence intervals of environments in stages (a) andbiozones (b) Sublittoral environments are always morediverse than nearshore environments but for theKarpatian (i.e. the Uvigerina gracliformis zone) thedifferences are not significant as indicated byoverlapping confidence intervals. Strong diversitydifferences between sublittoral assemblages at the levelof stages and biozones are evident, most notably betweenthe well-sampled Uvigerina graciliformis zone of theKarpatian and the Lower and Upper Lagenidae zones ofthe Badenian and point to habitat differences of therespective assemblages. K ns ¼ Karpatian nearshore;B ns ¼ Badenian nearshore; K sl ¼ Karpatian sublittoral;B sl ¼ Badenian sublittoral; LLZ ns ¼ Lower Lagenidaezone nearshore; LLZ sl ¼ Lower Lagenidae zonesublittoral; ULZ ns ¼ Upper Lagenidae zone nearshore;ULZ sl ¼ Upper Lagenidae zone sublittoral;BBZ ns ¼ Bolivina/Bulimina zone nearshore;BBZ sl ¼ Bolivina/Bulimina zone sublittoral;UgZ ns ¼ Uvigerina graciliformis zone nearshore;UgZ sl ¼ Uvigerina graciliformis zone sublittoral.

Fig. 11. Non-metric multidimensional scaling (nMDS)of family level data of the studied assemblages suggeststhe presence of a distinct water depth gradient alongaxis 1. Samples from nearshore environments of allbiozones cluster at the left. Sublittoral samples from theUvigerina graciliformis zone of the Karpatian representshallower environments than those from the Lower andUpper Lagenidae zones. Differences between the latterare tentatively explained by substrate differences(samples from the Lower Lagenidae zone tend to be fromsandy environments, samples Upper Lagenidae zone arerather from pelitic environments). LLZ ¼ LowerLagenidae zone; ULZ ¼ Upper Lagenidae zone;BBZ ¼ Bolivina/Bulimina zone; UgZ ¼ Uvigerinagraciliformis zone. Numbers 1–8 in the plot refer tosome outliers. 1–4 are characterized by high abundancesof otherwise rare taxa. In 5–7 the number of specimens isrelatively low, taxonomic composition heterogeneousand environmental affiliation therefore not straightforward. 8 is a sample with very high number of shells,which are strongly dominated by one taxon.


levels with highly resolved stratigraphic control(e.g. Johnson & Curry 2001; Hendy et al. 2009).Knowledge of local abundances of organismsenables determination of sampling completeness(Koch 1987) and to recognize ecological reorganiz-ation of regional biota, which can be independentfrom standing diversity (Jackson et al. 1999; Toddet al. 2002). In line with these evidences, thispaper highlights the sheer sampling effort that isrequired to reach the asymptote of the collector’scurve (Figs 4, 5, 9 & 10), a feature that is wellknown from modern and fossil molluscan assem-blages (e.g. Jackson et al. 1999; Bouchet et al.2002; Zuschin & Oliver 2005) and which suggeststhat most species are rare (Gaston 1994; Harnik2009). The use of abundance data allowed therecognition of ecological changes across a stageboundary, which drive the observed diversityincrease and which can be explained by a strongenvironmental shift.

Environmental bias in stages and biozones

This study demonstrates strong differences in quan-titative molluscan composition between two suc-ceeding stages (Fig. 6), but it also underlines apredominance of nearshore and shallow sublittoralhabitats in the studied Karpatian versus a predomi-nance of somewhat deeper environments in thestudied Badenian outcrops. Since shelf environ-ments have a higher diversity than the physicallystressed nearshore environments, the diversityincrease from the Karpatian to the Badenian in our

dataset can be largely related to an environmentalshift. When considering biozones, this environ-mental shift is especially pronounced at the 3rdorder sequence boundary between the KarpatianUvigerina graciliformis and the Badenian LowerLagenidae zones. This pattern amplifies theimpression of a diversity increase due to the Lan-ghian transgression from a literal reading of thefossil record (Fig. 8). Following Jablonski (1980),it is therefore important to sample a single habitator across a suite of habitats when evaluating diver-sity changes through time.

Although our data are from a relatively smallsubset of the Central Paratethys, they are consideredas representative because a study on echinodermsfrom the whole Central Paratethys also showedthat comparable habitats of the Karpatian and Bade-nian stages had very similar faunas and diversities(Kroh 2007). This author specifically stressed thatthe non-presence of Karpatian shallow-water car-bonates in the rock record explains much of thelower echinoderm diversity compared to the Bade-nian. Our study adds a new aspect in demonstrat-ing that also among siliciclastic sediments a faciesshift from nearshore and shallow sublittoral habitatsin the Karpatian to somewhat deeper environmentsin the Badenian is responsible for diversitydifferences.

Palaeogeography and palaeoclimate

It may be possible that for palaeogeographicalreasons the non-preserved deeper shelf assemblagesof the Karpatian were less diverse than their pre-served Badenian counterparts. During the Karpatiana marine connection of the Central Paratethysexisted only with the Mediterranean Basin, via theSlovenian ‘Trans-Tethyan Trench corridor’ (Bis-tricic & Jenko 1985). In the Badenian, open connec-tions with the Eastern Paratethys may also haveexisted, although the timing of the connections ishighly controversial (Rogl 1998; Studencka et al.1998; Steininger & Wessely 2000; Popov et al.2004). In both time slices, however, the Mediterra-nean Basin was at least temporarily connectedto the Indo-Pacific, enabling water circulationbetween both oceans, although faunas differedconsiderably (Harzhauser et al. 2007). A palaeogeo-graphical scenario for the observed diversity differ-ences is therefore rather speculative and notsupported by hard data. From a palaeoclimatologi-cal perspective the differences between the timeslices are rather small. This is because the Karpatianand Lower to Middle Badenian were characterizedby subtropical temperatures of the Middle Mioceneclimate optimum (Bohme 2003; Latal et al. 2006;Bruch et al. 2007; Kern et al. 2010), whichenabled the presence of thermophilic molluscs at

Fig. 12. Area and environments of Karpatian andBadenian outcrops in Eastern Austria (modified afterKroh 2007). Badenian outcrops mostly preserve fullymarine environments. Karpatian terrestrial, fluvial,fluvio-marine and limnic environments are veryprominent, suggesting that most fossiliferous marineoutcrops in this stage are from nearshore environments.


that time in the Paratethyan Basins (Harzhauseret al. 2003). In fact, nearshore assemblages, whichare available from both time slices, do not supportthe scenario of higher Badenian diversities(Fig. 10a).

The sequence stratigraphic framework

In our study on 3rd order cycles from the CentralParatethys, most outcrops are from highstandsystems tracts (Fig. 2, Table 1). These are internallycharacterized by relatively gradual biofacies repla-cements with major faunal turnovers occurringat sequence boundaries (Zuschin et al. 2007), apattern that corresponds to sequence stratigraphicexpectations (e.g. Brett 1995, 1998; Holland2000). The dominance of HSTs corresponds wellto the fact that the thickest parts of the sedimentaryrecord were built at times of progradation and thatthe transgressive phases are only represented bythin levels (e.g. Jablonski 1980; Fursich et al.1991; Clifton 2006).

Among the studied sequences, however, diversi-ties clearly depend on facies (Figs 9 & 10), whichdiffer in a systematic way due to a biased sedi-mentary record. Karpatian shell beds are mostlypreserved from nearshore and shallow sublittoralenvironments, which discordantly overlay macro-fossil-poor Karpatian offshore clays, whereas fromthe Badenian mostly somewhat deeper shelf assem-blages are recorded. This is most evident in theLower Lagenidae zone, which completely lacksnearshore assemblages (Figs 8, 10 & 11).

But also later in the Badenian, nearshoreassemblages are strikingly underrepresented whencompared to the Karpatian (Fig. 8). Sequence strat-igraphic models predict that nearshore sediments ofthe HST will be eroded during subsequent 3rd ordersea-level drops. This would well explain the paucityof nearshore sediments in the three Badenian 3rdorder cycles. This interpretation is supported by3-D seismic reflection data, which reveal significantdrops of relative sea-level (90–120 m) between thecycles (Strauss et al. 2006). The dominance of suchenvironments and corresponding lack of somewhatdeeper water shelf assemblages in the Karpatian iscounterintuitive, however, and is probably relatedto the strong tectonic reorganization of the CentralParatethys at the Karpatian/Badenian boundary(Adamek et al. 2003). One explanation for the scar-ceness of shelf environments is the uplift of theNorth Alpine Foreland Basin and the subsequentretreat of the sea. Deeper marine environmentsbecame established only in the Carpathian Foredeep(Rogl 1998). In contrast, the new tectonic regimeinitiated rapid subsidence in small satellite basinsof the Vienna Basin, where such littoral depositsescaped erosion (Wessely 1998; Kern et al. 2010).

Tectonics therefore affected sequence architecturein this particular setting by controlling subsidenceand sedimentary input, highlighting the problemthat sequence stratigraphic models were conceivedfor passive margin and only poorly predict sedimentaccumulation in tectonically active settings.

Comparison with other studies

Many studies have treated the distribution and pres-ervation of shell beds in relation to flooding surfacesand sequence boundaries (e.g. Kidwell 1988, 1989,1991; Banarjee & Kidwell 1991; Abbott & Carter1997; Kondo et al. 1998; Fursich & Pandey 2003).A series of others have examined palaeocommunitydynamics at local to regional scales in relation to therock record (e.g. Patzkowsky & Holland 1999;Goldman et al. 1999; Olszewski & Patzkowsky2003; Olszewski & Erwin 2004; Scarponi & Kowa-lewski 2004; Hendy & Kamp 2004; Dominici &Kowalke 2007; Zuschin et al. 2007; Tomasovych& Siblık 2007). Only few studies, however, havedealt with diversity changes as related to deposi-tional sequences. The results depend on scale,tectonic setting and environments preserved (oravailable to sample). Diversity seems largely to bedecoupled from 1st order cycles because stage-levelpost-Palaeozoic marine standing diversity ofwestern Europe increases although marine sedimentoutcrop area decreases (Smith 2001; see also Smith& McGowan 2007). A strong relation, however, hasbeen proposed for 2nd order sequence stratigraphiccycles (Smith 2001). Two case studies suggesthighest diversity or sampling probability at mid-cycle position at the top of transgressive systemstract intervals (Smith et al. 2001; Crampton et al.2006), but the causes seem to differ somewhatbetween tectonic settings (see discussion in Cramp-ton et al. 2006). At the active margin of NewZealand, for example, the best preservation of mol-luscan faunas is at mid-cycle position at the top oftransgressive systems tracts, and poorest preser-vation towards the end of highstand systems tracts.This is related to continuous subsidence and cre-ation of accommodation space (Crampton et al.2006). At the passive margin of western Europe,due to minimum accommodation space, shallow-water deposition is displaced onto the cratonicinteriors, where erosive loss during subsequentlowstands is most pervasive (Smith et al. 2001). Inboth areas, however, long-term diversity trendsare related to distinct facies biases. In theCenomanian/Turonian of western Europe a distinctdiversity decrease can be related to an increase ofoffshore at the expense of onshore sedimentaryfacies in the course of platform drowning due tosea-level rise (Smith et al. 2001). In the Neogeneof New Zealand an apparent decline in species


diversity reflects erosion of shallow-water depositsand a relative increase of bathyal at the cost ofshelf facies (Crampton et al. 2003). The importanceof environments covered within systems tracts isfinally also stressed in a study on late Quaternary4th order sequences deposited on the Po Plain(Italy). There, transgressive systems tract samplesdisplayed the highest, and the highstand systemstract samples the lowest diversity. At the sametime, turnover across sequences is negligible andmajor diversity shifts across systems tracts aremostly driven by Waltherian-type environmentalshifts (Scarponi & Kowalewski 2007).

Conclusions

The diversity increase between two regional stagesof the Central Paratethys is largely due to anenvironmental shift, which is related to selectivepreservation and erosion of environments due to tec-tonics and sea-level drops. Although most samplesanalysed in this study stem from highstand systemstracts, diversity differences between stages and bio-zones are significant. Pure standing diversity esti-mates will reveal biogeographical relations andmight capture faunal migrations aside from reflect-ing palaeoecological and palaeoclimatic bench-marks. They will not, however, reliably mirrorbiodiversity. This study therefore strongly supportsthe importance of environmental bias when consid-ering faunal changes though time and suggests thatin second and higher order sequences the faciescovered within systems tracts will drive diversitypatterns. The importance of rigorous, extensivesampling within a highly resolved stratigraphicand palaeoenvironmental framework for decipher-ing palaeodiversity patterns at regional scales isemphasized. The sheer sampling effort that isrequired to reach the asymptote of the collector’scurve is highlighted and it is strongly recommendedto use abundance data, which enable the recognitionof ecological changes in regional biota.

We thank S. Coric, P. Pervesler, R. Roetzel and numer-ous students for help with sampling; S. Dominici,J. Hohenegger, A. Kroh, F. Rogl, J. Sawyer and A. J.Tomasovych for discussions; A. Kroh and J. Reischer forproviding the data for Figure 12; A. Smith and A. J.McGowan for letting us present this work at the Lyell sym-posium 2010; A. J. McGowan and J. Nebelsick fora stimulating reviews of the manuscript. This studywas supported by project P19013-B17 of the AustrianScience Fund (FWF).

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